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<document id="6283F6F2927CAEBF7BE4815EAEEA46B5" ID-CLB-Dataset="7657" ID-DOI="10.1093/zoolinnean/zlz122" ID-GBIF-Dataset="ce6eb74d-6590-47d5-ba3a-a730db9a3baa" ID-Zenodo-Dep="4623663" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1616170722525" checkinUser="valdenar" docAuthor="Møbjerg, Nadja, Jørgensen, Aslak &amp; Kristensen, Reinhardt M." docDate="2020" docId="AA47465C3D3FFF892992FB42FE095874" docLanguage="en" docName="ZoolJLinnSoc.188.3.663-680.pdf" docOrigin="Zoological Journal of the Linnean Society 188 (3)" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-2023.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-2023.journal_article" docStyleVersion="7" docTitle="Anisonychidae MOBJERG, JORGENSEN &amp; KRISTENSEN 2020, FAM. NOV." docType="treatment" docVersion="12" lastPageNumber="666" masterDocId="567E3E243D3CFF8C2921FFF4FFF35943" masterDocTitle="Ongoing revision of Echiniscoididae (Heterotardigrada: Echiniscoidea), with the description of a new interstitial species and genus with unique anal structures" masterLastPageNumber="680" masterPageNumber="663" pageNumber="666" updateTime="1698934381141" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
<mods:mods id="C4FAE7CE982A76274F6B57A343E12C56" xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:title id="7CDED21E83DAC5BD2C9D8A5092567813">Ongoing revision of Echiniscoididae (Heterotardigrada: Echiniscoidea), with the description of a new interstitial species and genus with unique anal structures</mods:title>
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<mods:name id="190094DB27D8D0E804CAD19F68C37DAC" type="personal">
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<mods:namePart id="9B049ABC141B7FC7C0FD2230E2F12066">Møbjerg, Nadja</mods:namePart>
<mods:affiliation id="81D69EC1A051ED5ECD2AF351812275B0">Department of Biology, August Krogh Building, University of Copenhagen, Denmark</mods:affiliation>
<mods:nameIdentifier id="EFDC9E6359B43E68A15062E56470C377" type="email">nmobjerg@bio.ku.dk</mods:nameIdentifier>
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<mods:namePart id="C059DC81FAA6B4C060F06F3193F8D45F">Jørgensen, Aslak</mods:namePart>
<mods:affiliation id="9186D3BDE4704F7D28E546E52239C064">Department of Biology, August Krogh Building, University of Copenhagen, Denmark</mods:affiliation>
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<mods:namePart id="1F6E5D48DE0280D93209FFC4996EBC59">Kristensen, Reinhardt M.</mods:namePart>
<mods:affiliation id="D801B5959F484BCD3B0D3FB10509DEA5">The Natural History Museum of Denmark, University of Copenhagen, Denmark</mods:affiliation>
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<mods:title id="9600DD8169D64657E8925DCA4A0E087E">Zoological Journal of the Linnean Society</mods:title>
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<mods:part id="B3803C3DE47A6FBC26D707E323AC847B">
<mods:date id="1BC22E83E30520B6B3FE48320EDB5651">2020</mods:date>
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<mods:number id="E429921AA2CA466BC29753640E8FA83C">2019-09-18</mods:number>
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<mods:number id="D401E0208C70B7A29F5C90869BF66540">188</mods:number>
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<mods:number id="0103280A7ED490F518261D8A483211CD">3</mods:number>
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<treatment id="AA47465C3D3FFF892992FB42FE095874" ID-DOI="http://doi.org/10.5281/zenodo.4623671" ID-GBIF-Taxon="180598445" ID-Zenodo-Dep="4623671" LSID="urn:lsid:plazi:treatment:AA47465C3D3FFF892992FB42FE095874" httpUri="http://treatment.plazi.org/id/AA47465C3D3FFF892992FB42FE095874" lastPageId="5" lastPageNumber="666" pageId="3" pageNumber="666">
<subSubSection id="6AF4A4C13D3FFF8F2992FB42FDAD5DAE" pageId="3" pageNumber="666" type="nomenclature">
<paragraph id="2251F74A3D3FFF8F2992FB42FDAD5DAE" blockId="3.[163,779,1167,1330]" pageId="3" pageNumber="666">
<heading id="791940263D3FFF8F2992FB42FDAD5DAE" allCaps="true" centered="true" fontSize="9" level="2" pageId="3" pageNumber="666" reason="5">
FAMILY
<taxonomicName id="E5EE8CC93D3FFF8F282EFB42FE2C5DAF" ID-CoL="9b6391fc-bdcf-42ea-a059-f94633cb14c9" authority="MOBJERG, JORGENSEN &amp; KRISTENSEN" authorityName="MOBJERG, JORGENSEN &amp; KRISTENSEN" authorityYear="2020" class="Heterotardigrada" family="Anisonychidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Arthrotardigrada" pageId="3" pageNumber="666" phylum="Tardigrada" rank="family" status="fam. nov.">
<emphasis id="109A2B583D3FFF8F282EFB42FE265D8F" bold="true" box="[271,469,1206,1230]" pageId="3" pageNumber="666">ANISONYCHIDAE</emphasis>
MØBJERG, JØRGENSEN &amp; KRISTENSEN
</taxonomicName>
<emphasis id="109A2B583D3FFF8F28C7FB2CFDAD5DAE" bold="true" box="[486,606,1237,1261]" pageId="3" pageNumber="666">
<taxonomicNameLabel id="0BA996233D3FFF8F28C7FB2CFDAD5DAE" box="[486,606,1237,1261]" pageId="3" pageNumber="666" rank="family">FAM. NOV.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="6AF4A4C13D3FFF8F2982FB09FE935C71" pageId="3" pageNumber="666" type="description">
<paragraph id="2251F74A3D3FFF8F2982FB09FE935C71" blockId="3.[163,779,1167,1330]" pageId="3" pageNumber="666">lsid:zoobank.org:act:6A303AC8-EFA1-4639-BDE8- CB5CB32813EB</paragraph>
</subSubSection>
<subSubSection id="6AF4A4C13D3FFF8F2982FA9FFB125837" pageId="3" pageNumber="666" type="diagnosis">
<paragraph id="2251F74A3D3FFF8F2982FA9FFB125837" blockId="3.[163,780,1387,1856]" lastBlockId="3.[827,1444,197,372]" pageId="3" pageNumber="666">
<emphasis id="109A2B583D3FFF8F2982FA9FFEB25CC1" box="[163,321,1387,1410]" italics="true" pageId="3" pageNumber="666">Diagnosis:</emphasis>
Unplated, interstitial, marine heterotardigrades. Four claws inserteddirectly on eachof the first three leg pairs, and three claws inserted directly on the fourth pair of legs (in adults). Paired basal hooks on all claws, with the two hooks oriented at right angles to one another. Median cephalic cirrus is always present, butcan bereduced. Cephalicappendagesincludeinternal and external cirri, lateral cirri (cirri A) and generally also primary clava and flat, often indistinct, secondary clava. Cirri Eis always present. Papillae on leg IV are often present, and sensory structures on legs I–III may also be present (presently seen in only one species,
<taxonomicName id="E5EE8CC93D3FFF8F2BA5F938FE1F5E62" authority="Grimaldi de Zio, D'Addabbo Gallo, Morone De Lucia &amp; D'Addabbo, 1987" authorityName="Grimaldi de Zio, D'Addabbo Gallo, Morone De Lucia &amp; D'Addabbo" authorityYear="1987" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="species" species="mauritianus">
<emphasis id="109A2B583D3FFF8F2BA5F938FEDD5E42" italics="true" pageId="3" pageNumber="666">Anisonyches mauritianus</emphasis>
<bibRefCitation id="467F8ABB3D3FFF8F2815F918FE1F5E62" author="Grimaldi de Zio S &amp; D'Addabbo Gallo M &amp; Morone De Lucia MR &amp; D'Addabbo L." journalOrPublisher="Bollettino de Zoologia" pageId="3" pageNumber="666" pagination="347 - 357" part="4" refId="ref10604" refString="Grimaldi de Zio S, D'Addabbo Gallo M, Morone De Lucia MR, D'Addabbo L. 1987. Marine Arthrotardigrada and Echiniscoidea (Tardigrada, Heterotardigrada) from the Indian Ocean. Bollettino de Zoologia 4: 347 - 357." title="Marine Arthrotardigrada and Echiniscoidea (Tardigrada, Heterotardigrada) from the Indian Ocean" type="journal article" year="1987">Grimaldi de Zio, D’Addabbo Gallo, Morone De Lucia &amp; D’Addabbo, 1987</bibRefCitation>
</taxonomicName>
). Black eyes are present. Stylets are long, with stylet sheaths; stylet supports are lacking. The three placoids are CaCO 3 incrusted. Paired seminal receptacles are present in females. The coiled receptacles open relatively distant from the six-lobed female gonopore. The anal system consists of two lateral lobes connected by a median zig-zag structure, with the anus formed by a small terminal pore.
</paragraph>
</subSubSection>
<subSubSection id="6AF4A4C13D3FFF8F2A1AFE69FA8F5B10" pageId="3" pageNumber="666" type="type_taxon">
<paragraph id="2251F74A3D3FFF8F2A1AFE69FB0F58F7" blockId="3.[827,1276,413,436]" box="[827,1276,413,436]" pageId="3" pageNumber="666">
<emphasis id="109A2B583D3FFF8F2A1AFE69FC3B58F7" box="[827,968,413,436]" italics="true" pageId="3" pageNumber="666">
<typeStatus id="FD5549E83D3FFF8F2A1AFE69FC8758F7" box="[827,884,413,436]" pageId="3" pageNumber="666">Type</typeStatus>
genus:
</emphasis>
<taxonomicName id="E5EE8CC93D3FFF8F2AEFFE6AFB0B58F7" ID-CoL="WF2" authority="Pollock, 1975" authorityName="Pollock" authorityYear="1975" box="[974,1272,414,436]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2AEFFE6AFBAA58F0" box="[974,1113,414,435]" italics="true" pageId="3" pageNumber="666">Anisonyches</emphasis>
<bibRefCitation id="467F8ABB3D3FFF8F2D41FE6AFB0B58F7" author="Pollock LW" box="[1120,1272,414,436]" journalOrPublisher="Cahiers de Biologie Marine" pageId="3" pageNumber="666" pagination="121 - 132" part="16" refId="ref11508" refString="Pollock LW. 1975. Observations on marine Heterotardigrada including a new genus from the western Atlantic Ocean. Cahiers de Biologie Marine 16: 121 - 132." title="Observations on marine Heterotardigrada including a new genus from the western Atlantic Ocean" type="journal article" year="1975">Pollock, 1975</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="2251F74A3D3FFF8F2A1AFE29FC095B50" blockId="3.[827,1417,477,531]" pageId="3" pageNumber="666">
<emphasis id="109A2B583D3FFF8F2A1AFE29FB6E58B6" box="[827,1181,477,501]" italics="true" pageId="3" pageNumber="666">Generic diagnosis (amended)</emphasis>
: The same diagnosis as for the family.
</paragraph>
<paragraph id="2251F74A3D3FFF8F2A1AFDC8FA8F5B10" blockId="3.[827,1404,572,595]" box="[827,1404,572,595]" pageId="3" pageNumber="666">
<emphasis id="109A2B583D3FFF8F2A1AFDC8FC265B10" box="[827,981,572,595]" italics="true" pageId="3" pageNumber="666">
<typeStatus id="FD5549E83D3FFF8F2A1AFDC8FC875B10" box="[827,884,572,595]" pageId="3" pageNumber="666">Type</typeStatus>
species:
</emphasis>
<taxonomicName id="E5EE8CC93D3FFF8F2AFAFDC9FA8B5B10" ID-CoL="5V3C8" authority="Pollock, 1975" authorityName="Pollock" authorityYear="1975" box="[987,1400,573,595]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="species" species="diakidius">
<emphasis id="109A2B583D3FFF8F2AFAFDC9FB2A5B11" box="[987,1241,573,594]" italics="true" pageId="3" pageNumber="666">Anisonyches diakidius</emphasis>
<bibRefCitation id="467F8ABB3D3FFF8F2DC1FDC9FA8B5B10" author="Pollock LW" box="[1248,1400,573,595]" journalOrPublisher="Cahiers de Biologie Marine" pageId="3" pageNumber="666" pagination="121 - 132" part="16" refId="ref11508" refString="Pollock LW. 1975. Observations on marine Heterotardigrada including a new genus from the western Atlantic Ocean. Cahiers de Biologie Marine 16: 121 - 132." title="Observations on marine Heterotardigrada including a new genus from the western Atlantic Ocean" type="journal article" year="1975">Pollock, 1975</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="6AF4A4C13D3FFF892A1AFD8FFE095874" lastPageId="5" lastPageNumber="668" pageId="3" pageNumber="666" type="discussion">
<paragraph id="2251F74A3D3FFF8F2A1AFD8FFC1F5A4E" blockId="3.[827,1443,635,782]" pageId="3" pageNumber="666">
<emphasis id="109A2B583D3FFF8F2A1AFD8FFB8E5BD0" box="[827,1149,635,659]" italics="true" pageId="3" pageNumber="666">Other species in the genus:</emphasis>
<taxonomicName id="E5EE8CC93D3FFF8F2DA2FD89FC0F5B93" authority="Grimaldi de Zio, D'Addabbo Gallo, Morone De Lucia &amp; D'Addabbo, 1987" authorityName="Grimaldi de Zio, D'Addabbo Gallo, Morone De Lucia &amp; D'Addabbo" authorityYear="1987" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="species" species="mauritianus">
<emphasis id="109A2B583D3FFF8F2DA2FD89FA505BD0" box="[1155,1443,637,659]" italics="true" pageId="3" pageNumber="666">Anisonyches mauritianus</emphasis>
<bibRefCitation id="467F8ABB3D3FFF8F2A1AFD68FC0F5B93" author="Grimaldi de Zio S &amp; D'Addabbo Gallo M &amp; Morone De Lucia MR &amp; D'Addabbo L." journalOrPublisher="Bollettino de Zoologia" pageId="3" pageNumber="666" pagination="347 - 357" part="4" refId="ref10604" refString="Grimaldi de Zio S, D'Addabbo Gallo M, Morone De Lucia MR, D'Addabbo L. 1987. Marine Arthrotardigrada and Echiniscoidea (Tardigrada, Heterotardigrada) from the Indian Ocean. Bollettino de Zoologia 4: 347 - 357." title="Marine Arthrotardigrada and Echiniscoidea (Tardigrada, Heterotardigrada) from the Indian Ocean" type="journal article" year="1987">Grimaldi de Zio, D’Addabbo Gallo, Morone De Lucia &amp; D’Addabbo, 1987</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="E5EE8CC93D3FFF8F2D29FD4EFC805BAD" authority="Chang &amp; Rho, 1998" authorityName="Chang &amp; Rho" authorityYear="1998" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="species" species="deliquus">
<emphasis id="109A2B583D3FFF8F2D29FD4EFB085B8C" box="[1032,1275,698,719]" italics="true" pageId="3" pageNumber="666">Anisonyches deliquus</emphasis>
<bibRefCitation id="467F8ABB3D3FFF8F2C23FD4EFC805BAD" author="Chang CY &amp; Rho HS" journalOrPublisher="Korean Journal of Systematic Zoology" pageId="3" pageNumber="666" pagination="91 - 98" part="14" refId="ref10308" refString="Chang CY, Rho HS. 1998. Two marine tardigrade species of genus Anisonyches (Heterotardigrada: Echiniscoididae) from Mindanao, the Philippines. Korean Journal of Systematic Zoology 14: 91 - 98." title="Two marine tardigrade species of genus Anisonyches (Heterotardigrada: Echiniscoididae) from Mindanao, the Philippines" type="journal article" year="1998">Chang &amp; Rho, 1998</bibRefCitation>
</taxonomicName>
; and
<taxonomicName id="E5EE8CC93D3FFF8F2A91FD2DFC165A4E" authority="Bartels, Fontoura &amp; Nelson, 2018" authorityName="Bartels, Fontoura &amp; Nelson" authorityYear="2018" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="species" species="eleutherensis">
<emphasis id="109A2B583D3FFF8F2A91FD2DFB275BAD" box="[944,1236,729,750]" italics="true" pageId="3" pageNumber="666">Anisonyches eleutherensis</emphasis>
<bibRefCitation id="467F8ABB3D3FFF8F2DFAFD2DFC165A4E" author="Bartels PJ &amp; Fontoura P &amp; Nelson DR" journalOrPublisher="Zootaxa" pageId="3" pageNumber="666" pagination="043 - 070" part="4420" refId="ref10174" refString="Bartels PJ, Fontoura P, Nelson DR. 2018. Marine tardigrades of the Bahamas with the description of two new species and updated keys to the species of Anisonyches and Archechiniscus. Zootaxa 4420: 043 - 070." title="Marine tardigrades of the Bahamas with the description of two new species and updated keys to the species of Anisonyches and Archechiniscus" type="journal volume" year="2018">Bartels, Fontoura &amp; Nelson, 2018</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="2251F74A3D3FFF892A1AFCC0FE095874" blockId="3.[827,1444,820,1855]" lastBlockId="5.[163,779,197,311]" lastPageId="5" lastPageNumber="668" pageId="3" pageNumber="666">
<emphasis id="109A2B583D3FFF8F2A1AFCC0FC4C5A08" box="[827,959,820,843]" italics="true" pageId="3" pageNumber="666">Discussion</emphasis>
: For the present study, we have examined three specimens of the
<typeStatus id="FD5549E83D3FFF8F2D43FCA0FB645A2A" box="[1122,1175,852,873]" pageId="3" pageNumber="666">type</typeStatus>
species,
<taxonomicName id="E5EE8CC93D3FFF8F2C2BFCA0FA505A2A" authorityName="Pollock" authorityYear="1975" box="[1290,1443,852,873]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="species" species="diakidius">
<emphasis id="109A2B583D3FFF8F2C2BFCA0FA505A2A" box="[1290,1443,852,873]" italics="true" pageId="3" pageNumber="666">A. diakidius</emphasis>
</taxonomicName>
(
<figureCitation id="BAD5EBCF3D3FFF8F2A65FC86FC4C5ACB" box="[836,959,882,904]" captionStart="Figure 1" captionStartId="4.[145,224,1545,1567]" captionTargetBox="[226,1343,195,1505]" captionTargetId="figure-28@4.[606,1343,621,1009]" captionTargetPageId="4" captionText="Figure 1. Light microscopy and habitus drawing of Anisonyches diakidius from Guadeloupe collected by Jeanne Renaud- Mornant, 1979 and deposited at the Natural History Museum of Denmark (ZMUC-TAR 1200–1202). A, lateral view of a female (ZMUC-TAR 1202) with indistinct median cirrus.Scale bar: 50 µm. B, close-upof seminal receptacle of the samefemale (ZMUC-TAR 1202).Lateral view.Scale bar:10 µm.C, close-upof head region (ZMUC-TAR 1202) revealing the buccal tube, the thin median cirrus and the pharyngeal bulb.Scale bar: 10 µm. D, ventral view (ZMUC-TAR 1201) of female gonopore and two of the claws of the fourth leg. The inset shows the claws of the second leg. Note the basal hook of the claws. Scale bars: 10 µm. E, dorsal view of female (ZMUC-TAR 1200) showing various sensory structures and claws. Scale bar: 50 µm. F, habitus drawing of female (ZMUC-TAR 1202) in dorsal position, before it moved into a lateral position, made by J. Renaud-Mornant in 1979, redrawn by S. Elle in 2019. Scale bar: 20 µm. Abbreviations: an, anus; bh, basal hook of claw; bt, buccal tube; cE, cirrus E; cl, claw; ec, external cirrus; go, gonopore; ic, internal cirrus; lc, lateral cirrus (cirrus A); Leg 4, terminal leg; mc, median cirrus; mg, midgut; p4, sensory papilla of fourth leg; pb, pharyngeal bulb; pc, primary clava; pl, placoid; sc, secondary clava, sg, stylet gland; sr, seminal receptacle; ss, stylet sheath; st, stylet." figureDoi="http://doi.org/10.5281/zenodo.4623669" httpUri="https://zenodo.org/record/4623669/files/figure.png" pageId="3" pageNumber="666">Fig. 1A–E</figureCitation>
). These specimens (ZMUC-TAR 1200– 1202) were collected by J. Renaud-Mornant in 1979 from Guadeloupe (French Antilles) and sent to our museum along with a drawing of one of the specimens (
<figureCitation id="BAD5EBCF3D3FFF8F2A62FC19FC605D41" box="[835,915,1005,1027]" captionStart="Figure 1" captionStartId="4.[145,224,1545,1567]" captionTargetBox="[226,1343,195,1505]" captionTargetId="figure-28@4.[606,1343,621,1009]" captionTargetPageId="4" captionText="Figure 1. Light microscopy and habitus drawing of Anisonyches diakidius from Guadeloupe collected by Jeanne Renaud- Mornant, 1979 and deposited at the Natural History Museum of Denmark (ZMUC-TAR 1200–1202). A, lateral view of a female (ZMUC-TAR 1202) with indistinct median cirrus.Scale bar: 50 µm. B, close-upof seminal receptacle of the samefemale (ZMUC-TAR 1202).Lateral view.Scale bar:10 µm.C, close-upof head region (ZMUC-TAR 1202) revealing the buccal tube, the thin median cirrus and the pharyngeal bulb.Scale bar: 10 µm. D, ventral view (ZMUC-TAR 1201) of female gonopore and two of the claws of the fourth leg. The inset shows the claws of the second leg. Note the basal hook of the claws. Scale bars: 10 µm. E, dorsal view of female (ZMUC-TAR 1200) showing various sensory structures and claws. Scale bar: 50 µm. F, habitus drawing of female (ZMUC-TAR 1202) in dorsal position, before it moved into a lateral position, made by J. Renaud-Mornant in 1979, redrawn by S. Elle in 2019. Scale bar: 20 µm. Abbreviations: an, anus; bh, basal hook of claw; bt, buccal tube; cE, cirrus E; cl, claw; ec, external cirrus; go, gonopore; ic, internal cirrus; lc, lateral cirrus (cirrus A); Leg 4, terminal leg; mc, median cirrus; mg, midgut; p4, sensory papilla of fourth leg; pb, pharyngeal bulb; pc, primary clava; pl, placoid; sc, secondary clava, sg, stylet gland; sr, seminal receptacle; ss, stylet sheath; st, stylet." figureDoi="http://doi.org/10.5281/zenodo.4623669" httpUri="https://zenodo.org/record/4623669/files/figure.png" pageId="3" pageNumber="666">Fig. 1F</figureCitation>
). Furthermore, we have examined a specimen of
<taxonomicName id="E5EE8CC93D3FFF8F2A79FBF8FBFE5D62" authorityName="Bartels, Fontoura &amp; Nelson" authorityYear="2018" box="[856,1037,1036,1057]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="species" species="eleutherensis">
<emphasis id="109A2B583D3FFF8F2A79FBF8FBFE5D62" box="[856,1037,1036,1057]" italics="true" pageId="3" pageNumber="666">A. eleutherensis</emphasis>
</taxonomicName>
from the Bahamas, present in our collection(ZMUC-TAR1204),withadistinct
<taxonomicName id="E5EE8CC93D3FFF8F2C3BFBDEFA515D7C" authorityName="Pollock" authorityYear="1975" box="[1306,1442,1066,1087]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2C3BFBDEFA515D7C" box="[1306,1442,1066,1087]" italics="true" pageId="3" pageNumber="666">Anisonyches</emphasis>
</taxonomicName>
claw configuration as described by
<bibRefCitation id="467F8ABB3D3FFF8F2DE0FBBDFA6F5D1C" author="Bartels PJ &amp; Fontoura P &amp; Nelson DR" box="[1217,1436,1097,1119]" journalOrPublisher="Zootaxa" pageId="3" pageNumber="666" pagination="043 - 070" part="4420" refId="ref10174" refString="Bartels PJ, Fontoura P, Nelson DR. 2018. Marine tardigrades of the Bahamas with the description of two new species and updated keys to the species of Anisonyches and Archechiniscus. Zootaxa 4420: 043 - 070." title="Marine tardigrades of the Bahamas with the description of two new species and updated keys to the species of Anisonyches and Archechiniscus" type="journal volume" year="2018">
Bartels
<emphasis id="109A2B583D3FFF8F2C3BFBBEFAA25D1D" box="[1306,1361,1097,1119]" italics="true" pageId="3" pageNumber="666">et al.</emphasis>
(2018)
</bibRefCitation>
. From our recent investigations, and that of colleagues (
<bibRefCitation id="467F8ABB3D3FFF8F2A62FB72FBEE5DDF" author="Bartels PJ &amp; Fontoura P &amp; Nelson DR" box="[835,1053,1158,1180]" journalOrPublisher="Zootaxa" pageId="3" pageNumber="666" pagination="043 - 070" part="4420" refId="ref10174" refString="Bartels PJ, Fontoura P, Nelson DR. 2018. Marine tardigrades of the Bahamas with the description of two new species and updated keys to the species of Anisonyches and Archechiniscus. Zootaxa 4420: 043 - 070." title="Marine tardigrades of the Bahamas with the description of two new species and updated keys to the species of Anisonyches and Archechiniscus" type="journal volume" year="2018">
Bartels
<emphasis id="109A2B583D3FFF8F2ABEFB73FC2B5DD8" box="[927,984,1158,1180]" italics="true" pageId="3" pageNumber="666">et al.</emphasis>
, 2018
</bibRefCitation>
), it is apparent that
<taxonomicName id="E5EE8CC93D3FFF8F2C31FB72FA6F5DD8" authorityName="Pollock" authorityYear="1975" box="[1296,1436,1158,1179]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2C31FB72FA6F5DD8" box="[1296,1436,1158,1179]" italics="true" pageId="3" pageNumber="666">Anisonyches</emphasis>
</taxonomicName>
, with its seminal receptacles (
<figureCitation id="BAD5EBCF3D3FFF8F2DB5FB51FB1B5DF9" box="[1172,1256,1189,1211]" captionStart="Figure 1" captionStartId="4.[145,224,1545,1567]" captionTargetBox="[226,1343,195,1505]" captionTargetId="figure-28@4.[606,1343,621,1009]" captionTargetPageId="4" captionText="Figure 1. Light microscopy and habitus drawing of Anisonyches diakidius from Guadeloupe collected by Jeanne Renaud- Mornant, 1979 and deposited at the Natural History Museum of Denmark (ZMUC-TAR 1200–1202). A, lateral view of a female (ZMUC-TAR 1202) with indistinct median cirrus.Scale bar: 50 µm. B, close-upof seminal receptacle of the samefemale (ZMUC-TAR 1202).Lateral view.Scale bar:10 µm.C, close-upof head region (ZMUC-TAR 1202) revealing the buccal tube, the thin median cirrus and the pharyngeal bulb.Scale bar: 10 µm. D, ventral view (ZMUC-TAR 1201) of female gonopore and two of the claws of the fourth leg. The inset shows the claws of the second leg. Note the basal hook of the claws. Scale bars: 10 µm. E, dorsal view of female (ZMUC-TAR 1200) showing various sensory structures and claws. Scale bar: 50 µm. F, habitus drawing of female (ZMUC-TAR 1202) in dorsal position, before it moved into a lateral position, made by J. Renaud-Mornant in 1979, redrawn by S. Elle in 2019. Scale bar: 20 µm. Abbreviations: an, anus; bh, basal hook of claw; bt, buccal tube; cE, cirrus E; cl, claw; ec, external cirrus; go, gonopore; ic, internal cirrus; lc, lateral cirrus (cirrus A); Leg 4, terminal leg; mc, median cirrus; mg, midgut; p4, sensory papilla of fourth leg; pb, pharyngeal bulb; pc, primary clava; pl, placoid; sc, secondary clava, sg, stylet gland; sr, seminal receptacle; ss, stylet sheath; st, stylet." figureDoi="http://doi.org/10.5281/zenodo.4623669" httpUri="https://zenodo.org/record/4623669/files/figure.png" pageId="3" pageNumber="666">Fig. 1B</figureCitation>
), median cirrus (
<figureCitation id="BAD5EBCF3D3FFF8F2A62FB30FC605D9A" box="[835,915,1220,1242]" captionStart="Figure 1" captionStartId="4.[145,224,1545,1567]" captionTargetBox="[226,1343,195,1505]" captionTargetId="figure-28@4.[606,1343,621,1009]" captionTargetPageId="4" captionText="Figure 1. Light microscopy and habitus drawing of Anisonyches diakidius from Guadeloupe collected by Jeanne Renaud- Mornant, 1979 and deposited at the Natural History Museum of Denmark (ZMUC-TAR 1200–1202). A, lateral view of a female (ZMUC-TAR 1202) with indistinct median cirrus.Scale bar: 50 µm. B, close-upof seminal receptacle of the samefemale (ZMUC-TAR 1202).Lateral view.Scale bar:10 µm.C, close-upof head region (ZMUC-TAR 1202) revealing the buccal tube, the thin median cirrus and the pharyngeal bulb.Scale bar: 10 µm. D, ventral view (ZMUC-TAR 1201) of female gonopore and two of the claws of the fourth leg. The inset shows the claws of the second leg. Note the basal hook of the claws. Scale bars: 10 µm. E, dorsal view of female (ZMUC-TAR 1200) showing various sensory structures and claws. Scale bar: 50 µm. F, habitus drawing of female (ZMUC-TAR 1202) in dorsal position, before it moved into a lateral position, made by J. Renaud-Mornant in 1979, redrawn by S. Elle in 2019. Scale bar: 20 µm. Abbreviations: an, anus; bh, basal hook of claw; bt, buccal tube; cE, cirrus E; cl, claw; ec, external cirrus; go, gonopore; ic, internal cirrus; lc, lateral cirrus (cirrus A); Leg 4, terminal leg; mc, median cirrus; mg, midgut; p4, sensory papilla of fourth leg; pb, pharyngeal bulb; pc, primary clava; pl, placoid; sc, secondary clava, sg, stylet gland; sr, seminal receptacle; ss, stylet sheath; st, stylet." figureDoi="http://doi.org/10.5281/zenodo.4623669" httpUri="https://zenodo.org/record/4623669/files/figure.png" pageId="3" pageNumber="666">Fig. 1C</figureCitation>
) and unique claw configuration (
<figureCitation id="BAD5EBCF3D3FFF8F2C2BFB30FAAF5D9A" box="[1290,1372,1220,1242]" captionStart="Figure 1" captionStartId="4.[145,224,1545,1567]" captionTargetBox="[226,1343,195,1505]" captionTargetId="figure-28@4.[606,1343,621,1009]" captionTargetPageId="4" captionText="Figure 1. Light microscopy and habitus drawing of Anisonyches diakidius from Guadeloupe collected by Jeanne Renaud- Mornant, 1979 and deposited at the Natural History Museum of Denmark (ZMUC-TAR 1200–1202). A, lateral view of a female (ZMUC-TAR 1202) with indistinct median cirrus.Scale bar: 50 µm. B, close-upof seminal receptacle of the samefemale (ZMUC-TAR 1202).Lateral view.Scale bar:10 µm.C, close-upof head region (ZMUC-TAR 1202) revealing the buccal tube, the thin median cirrus and the pharyngeal bulb.Scale bar: 10 µm. D, ventral view (ZMUC-TAR 1201) of female gonopore and two of the claws of the fourth leg. The inset shows the claws of the second leg. Note the basal hook of the claws. Scale bars: 10 µm. E, dorsal view of female (ZMUC-TAR 1200) showing various sensory structures and claws. Scale bar: 50 µm. F, habitus drawing of female (ZMUC-TAR 1202) in dorsal position, before it moved into a lateral position, made by J. Renaud-Mornant in 1979, redrawn by S. Elle in 2019. Scale bar: 20 µm. Abbreviations: an, anus; bh, basal hook of claw; bt, buccal tube; cE, cirrus E; cl, claw; ec, external cirrus; go, gonopore; ic, internal cirrus; lc, lateral cirrus (cirrus A); Leg 4, terminal leg; mc, median cirrus; mg, midgut; p4, sensory papilla of fourth leg; pb, pharyngeal bulb; pc, primary clava; pl, placoid; sc, secondary clava, sg, stylet gland; sr, seminal receptacle; ss, stylet sheath; st, stylet." figureDoi="http://doi.org/10.5281/zenodo.4623669" httpUri="https://zenodo.org/record/4623669/files/figure.png" pageId="3" pageNumber="666">Fig. 1D</figureCitation>
), does not belong among the echiniscoidids. Consequently, based on the presence of a median cephalic cirrus and seminal receptacles, we place
<taxonomicName id="E5EE8CC93D3FFF8F2DB8FAD4FAD55C76" authorityName="Pollock" authorityYear="1975" box="[1177,1318,1312,1333]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2DB8FAD4FAD55C76" box="[1177,1318,1312,1333]" italics="true" pageId="3" pageNumber="666">Anisonyches</emphasis>
</taxonomicName>
in its own family,
<taxonomicName id="E5EE8CC93D3FFF8F2AAFFACAFBCB5C17" authorityName="MOBJERG, JORGENSEN &amp;" authorityYear="2020" box="[910,1080,1342,1364]" class="Heterotardigrada" family="Anisonychidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Arthrotardigrada" pageId="3" pageNumber="666" phylum="Tardigrada" rank="family">Anisonychidae</taxonomicName>
, in the order
<taxonomicName id="E5EE8CC93D3FFF8F2DF5FACAFA6C5C17" authorityName="Marcus" authorityYear="1927" box="[1236,1439,1342,1364]" class="Heterotardigrada" higherTaxonomySource="GBIF" kingdom="Animalia" order="Arthrotardigrada" pageId="3" pageNumber="666" phylum="Tardigrada" rank="order">Arthrotardigrada</taxonomicName>
, but at the same time strongly emphasize that
<taxonomicName id="E5EE8CC93D3FFF8F2A1AFA88FBF65CD1" authorityName="Marcus" authorityYear="1927" box="[827,1029,1404,1426]" class="Heterotardigrada" higherTaxonomySource="GBIF" kingdom="Animalia" order="Arthrotardigrada" pageId="3" pageNumber="666" phylum="Tardigrada" rank="order">Arthrotardigrada</taxonomicName>
is paraphyletic and thus in need of taxonomic revision. We note that the claw configuration of
<taxonomicName id="E5EE8CC93D3FFF8F2A76FA4DFC175C8D" authorityName="Pollock" authorityYear="1975" box="[855,996,1465,1486]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2A76FA4DFC175C8D" box="[855,996,1465,1486]" italics="true" pageId="3" pageNumber="666">Anisonyches</emphasis>
</taxonomicName>
bears some resemblance to the limnic
<taxonomicName id="E5EE8CC93D3FFF8F2A1AFA2CFAD45CAD" authority="Binda &amp; Kristensen, 1986" authorityName="Binda &amp; Kristensen" authorityYear="1986" box="[827,1319,1496,1518]" class="Heterotardigrada" family="Carphanidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="family">
Carphanidae
<bibRefCitation id="467F8ABB3D3FFF8F2AC2FA2CFAD45CAD" author="Binda MG &amp; Kristensen RM" box="[995,1319,1496,1518]" journalOrPublisher="Animalia" pageId="3" pageNumber="666" pagination="9 - 20" part="13" refId="ref10265" refString="Binda MG, Kristensen RM. 1986. Notes on the genus Oreella (Oreellidae) and the systematic position of Carphania fluviatilis Binda, 1978 (Carphanidae fam. nov., Heterotardigrada). Animalia 13: 9 - 20." title="Notes on the genus Oreella (Oreellidae) and the systematic position of Carphania fluviatilis Binda, 1978 (Carphanidae fam. nov., Heterotardigrada)" type="journal article" year="1986">Binda &amp; Kristensen, 1986</bibRefCitation>
</taxonomicName>
currently retained in
<taxonomicName id="E5EE8CC93D3FFF8F2AE4FA02FB9A5F4F" authorityName="Richters" authorityYear="1926" box="[965,1129,1526,1548]" class="Heterotardigrada" higherTaxonomySource="GBIF" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="order">Echiniscoidea</taxonomicName>
. Within
<taxonomicName id="E5EE8CC93D3FFF8F2DECFA02FA835F4F" authorityName="Richters" authorityYear="1926" box="[1229,1392,1526,1548]" class="Heterotardigrada" higherTaxonomySource="GBIF" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="order">Echiniscoidea</taxonomicName>
, the family
<taxonomicName id="E5EE8CC93D3FFF8F2AAAF9E1FBF15F68" authorityName="Ramazzotti" authorityYear="1962" box="[907,1026,1557,1579]" class="Heterotardigrada" family="Oreellidae" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="family">Oreellidae</taxonomicName>
has retained seminal receptacles as in
<taxonomicName id="E5EE8CC93D3FFF8F2A79F9C0FC135F0A" authorityName="Pollock" authorityYear="1975" box="[856,992,1588,1609]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2A79F9C0FC135F0A" box="[856,992,1588,1609]" italics="true" pageId="3" pageNumber="666">Anisonyches</emphasis>
</taxonomicName>
, but the claw configuration in
<taxonomicName id="E5EE8CC93D3FFF8F2C19F9C0FA7A5F0A" authorityName="Murray" authorityYear="1910" box="[1336,1417,1588,1609]" class="Heterotardigrada" family="Oreellidae" genus="Oreella" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2C19F9C0FA7A5F0A" box="[1336,1417,1588,1609]" italics="true" pageId="3" pageNumber="666">Oreella</emphasis>
</taxonomicName>
is different from
<taxonomicName id="E5EE8CC93D3FFF8F2AFFF9A6FB945F24" authorityName="Pollock" authorityYear="1975" box="[990,1127,1618,1639]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2AFFF9A6FB945F24" box="[990,1127,1618,1639]" italics="true" pageId="3" pageNumber="666">Anisonyches</emphasis>
</taxonomicName>
and
<taxonomicName id="E5EE8CC93D3FFF8F2DBFF9A6FAEB5F24" authorityName="Binda" authorityYear="1978" box="[1182,1304,1618,1639]" class="Heterotardigrada" family="Carphaniidae" genus="Carphania" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2DBFF9A6FAEB5F24" box="[1182,1304,1618,1639]" italics="true" pageId="3" pageNumber="666">Carphania</emphasis>
</taxonomicName>
with regard to the position of the basal hooks, putative indications of toes and four claws on all legs (
<bibRefCitation id="467F8ABB3D3FFF8F2D9BF964FC825F87" author="Binda MG &amp; Kristensen RM" journalOrPublisher="Animalia" pageId="3" pageNumber="666" pagination="9 - 20" part="13" refId="ref10265" refString="Binda MG, Kristensen RM. 1986. Notes on the genus Oreella (Oreellidae) and the systematic position of Carphania fluviatilis Binda, 1978 (Carphanidae fam. nov., Heterotardigrada). Animalia 13: 9 - 20." title="Notes on the genus Oreella (Oreellidae) and the systematic position of Carphania fluviatilis Binda, 1978 (Carphanidae fam. nov., Heterotardigrada)" type="journal article" year="1986">Binda &amp; Kristensen, 1986</bibRefCitation>
). Here, we emphasize the fact that both caphanids and oreellids lack a median cirrus.
<bibRefCitation id="467F8ABB3D3FFF8F2DC7F939FC4C5E41" author="Grimaldi de Zio S &amp; D'Addabbo Gallo M &amp; Morone De Lucia MR &amp; D'Addabbo L." journalOrPublisher="Bollettino de Zoologia" pageId="3" pageNumber="666" pagination="347 - 357" part="4" refId="ref10604" refString="Grimaldi de Zio S, D'Addabbo Gallo M, Morone De Lucia MR, D'Addabbo L. 1987. Marine Arthrotardigrada and Echiniscoidea (Tardigrada, Heterotardigrada) from the Indian Ocean. Bollettino de Zoologia 4: 347 - 357." title="Marine Arthrotardigrada and Echiniscoidea (Tardigrada, Heterotardigrada) from the Indian Ocean" type="journal article" year="1987">
Grimaldi de Zio
<emphasis id="109A2B583D3FFF8F2A1AF918FC815E42" box="[827,882,1772,1793]" italics="true" pageId="3" pageNumber="666">et al.</emphasis>
(1987)
</bibRefCitation>
suggested that
<taxonomicName id="E5EE8CC93D3FFF8F2D51F918FB0B5E42" authorityName="Pollock" authorityYear="1975" box="[1136,1272,1772,1793]" class="Heterotardigrada" family="Echiniscoididae" genus="Anisonyches" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="genus">
<emphasis id="109A2B583D3FFF8F2D51F918FB0B5E42" box="[1136,1272,1772,1793]" italics="true" pageId="3" pageNumber="666">Anisonyches</emphasis>
</taxonomicName>
is more closely related to the arthrotardigrade taxa
<taxonomicName id="E5EE8CC93D3FFF8F2DC3F8FEFA765E63" authorityName="Renaud-Mornant" authorityYear="1974" box="[1250,1413,1802,1824]" class="Heterotardigrada" family="Coronarctidae" kingdom="Animalia" order="Arthrotardigrada" pageId="3" pageNumber="666" phylum="Tardigrada" rank="family">Coronarctidae</taxonomicName>
or
<taxonomicName id="E5EE8CC93D3FFF8F2A1AF8DDFC3A5E7C" authorityName="Schulz" authorityYear="1951" box="[827,969,1833,1855]" class="Heterotardigrada" family="Stygarctidae" kingdom="Animalia" order="Arthrotardigrada" pageId="3" pageNumber="666" phylum="Tardigrada" rank="family">Stygarctidae</taxonomicName>
thanto
<taxonomicName id="E5EE8CC93D3FFF8F2D3EF8DDFB225E7C" baseAuthorityName="Mobjerg et al." baseAuthorityYear="2016" box="[1055,1233,1833,1855]" class="Heterotardigrada" family="Echiniscoididae" kingdom="Animalia" order="Echiniscoidea" pageId="3" pageNumber="666" phylum="Tardigrada" rank="family">Echiniscoididae</taxonomicName>
. Wenote that claws inserted directly on the leg is a characteristic of both
<taxonomicName id="E5EE8CC93D39FF892982FF10FEBA59B9" authorityName="Renaud-Mornant" authorityYear="1974" box="[163,329,228,250]" class="Heterotardigrada" family="Coronarctidae" kingdom="Animalia" order="Arthrotardigrada" pageId="5" pageNumber="668" phylum="Tardigrada" rank="family">Coronarctidae</taxonomicName>
and
<taxonomicName id="E5EE8CC93D39FF8928A7FF10FDEF59B9" authorityName="Schulz" authorityYear="1951" box="[390,540,228,250]" class="Heterotardigrada" family="Stygarctidae" kingdom="Animalia" order="Arthrotardigrada" pageId="5" pageNumber="668" phylum="Tardigrada" rank="family">Stygarctidae</taxonomicName>
, clearly underlining the need for thorough future investigations into limnic and marine heterotardigrades.
</paragraph>
</subSubSection>
</treatment>
</document>
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